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Lucidi, P., Bacco, G., Sticco, M., Mazzoleni, G., Benvenuti, M., Bernabò, N., et al. (2013). Assessment of motor laterality in foals and young horses (Equus caballus) through an analysis of derailment at trot. Physiol. Behav., 109, 8–13.
Abstract: The conflicting results regarding the study of motor laterality in horses may indicate that there does not exist a proper method to assess the degree and the direction of motor bias in these animals. Unfortunately, even less is known about the development of laterality in horses, and to what extent early manipulations can still exert their effects in adulthood. We propose a new method that can be easily applied at a very early age thus avoiding testing adult horses eventually biased by human handling and/or training. Forty-six horses (29 nine-month-old foals and 17 two-year old horses) were handled since birth bilaterally and housed in groups in wide areas. At the time of the analysis, in order to minimize environmental and sensorial disturbances, each horse was tested in a round pen individually or as dyad mother-foal. The ability/inability to properly execute a circle at trot was then recorded, assuming the direction of derailment, i.e. the cutting of the circle, as an indicator of motor bias. From the results of the study it is arguable that motor laterality in horses is acquired over time: in fact foals tested while their mothers were being subjected to longeing showed a higher percentage of ambidextrous animals, while two-year-old horses appeared biased toward the right (p<0.05). Results are discussed in the light of the scientific knowledge about equine biomechanics, taking into account horses' locomotion that leads to the advancement of the body mass through the activation of a kinetic chain that originates from the hindquarters.
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Yarnell, K., Hall, C., & Billett, E. (2013). An assessment of the aversive nature of an animal management procedure (clipping) using behavioral and physiological measures. Physiol. Behav., 118, 32–39.
Abstract: Animal management often involves procedures that, while unlikely to cause physical pain, still cause aversive responses. The domestic horse (Equus caballus) regularly has excessive hair clipped off to facilitate its use as a riding/driving animal and this procedure causes adverse behavioral responses in some animals. The aim of this study was to compare behavioral and physiological measures to assess the aversive effect of this procedure. Ten horses were selected on the basis of being either compliant (C: n=5) or non-compliant (NC: n=5) during this procedure. The horses were subjected to a sham clipping procedure (SC: where the blades had been removed from the clippers) for a period of ten minutes. Measures were taken pre, during and post SC (-10min to +30min) and mean values calculated for ALL horses and for C and NC separately. Behavioral activity was scored (scale 1-5) by twenty students from video footage in (phase/group-blind scoring). Heart rate (HR), salivary cortisol and eye temperature were monitored throughout the procedure. The NC horses were found to be significantly more behaviorally active/less relaxed throughout the trial than C horses (p<0.05) with the greatest difference occurring during the SC procedure (p<0.01). NC horses were more active/less relaxed during, compared with pre or post SC (p<0.05), but showed no behavioral difference pre and post SC. HR of the NC horses was higher than that of the C horses throughout the trial but only significantly so after 10min of SC (p<0.01). ALL horses showed a significant increase in HR between +5 and +10min into the procedure (p<0.05). There was a significant increase in salivary cortisol concentration in ALL horses post procedure (p<0.01) with levels peaking at 20minute post SC. No significant differences in salivary cortisol concentration between C and NC were found at any stage of the trial. Eye temperature increased significantly in ALL horses during SC, peaking at +10min into the procedure (p<0.05) and then decreased substantially when SC had ceased (p<0.01). Although no significant differences were found between C and NC per se, there was a significant interaction between group and phase of trial (p<0.05) with the NC group showing a greater decrease in eye temperature post SC. There was a significant positive correlation between changes in salivary cortisol concentration and eye temperature (p<0.01) but no correlation between any of the other measures. Although the behavioral response of C and NC to this procedure was significantly different the physiological responses indicated that ALL horses found the procedure aversive. Eye temperature could be used as an objective and immediate measure of how an animal is responding to a specific situation in order to evaluate management procedures and adapt them where appropriate to reduce the negative impact on animal health and welfare.
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Palme, R. (2019). Non-invasive measurement of glucocorticoids: Advances and problems. Physiol. Behav., 199, 229–243.
Abstract: Glucocorticoids (GCs; i.e. cortisol/corticosterone) are a central component of the stress response and thus their measurement is frequently used to evaluate the impact of stressful situations. Their metabolites from faeces of various animal species are more and more taken as a non-invasive aid to assess GC release and thus adrenocortical activity. The current literature review includes an extensive collection (1327 papers) and evaluation (see also Supplementary Tables) of the literature on faecal cortisol/corticosterone metabolite (FCM) analysis published to date. It aims at giving reference for researchers interested in implementing FCM analysis into their study or seeking to improve such methods by providing background knowledge on GC metabolism and excretion, conveying insights into methodological issues and stating caveats of FCM analysis and by highlighting prerequisites for and some examples of a successful application of such methods. Collecting faecal samples and analysing FCMs may appear simple and straightforward, but researchers have to select and apply methods correctly. They also need to be aware of the many pitfalls and potentially confounding factors and, last but not least, have to carefully interpret results. Applied properly, measurement of FCMs is a powerful non-invasive tool in a variety of research areas, such as (stress) biology, ethology, ecology, animal conservation and welfare, but also biomedicine.
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Christensen, J. W., Beekmans, M., van Dalum, M., & VanDierendonck, M. (2014). Effects of hyperflexion on acute stress responses in ridden dressage horses. Physiol. Behav., 128, 39–45.
Abstract: The effects of hyperflexion on the welfare of dressage horses have been debated. This study aimed to investigate acute stress responses of dressage horses ridden in three different Head-and-Neck-positions (HNPs). Fifteen dressage horses were ridden by their usual rider in a standardised 10-min dressage programme in either the competition frame (CF), hyperflexion (“Low-Deep-and-Round”; LDR) or a looser frame (LF) in a balanced order on three separate test days. Heart rate (HR), heart rate variability parameters (HRV), behaviour and rein tension were recorded during the test. Salivary cortisol concentrations were measured 60min before and 0, 5, 15 and 30min after the test. Rein tension was significantly lower in LF and did not differ between CF and LDR; however approx. 15% of recordings in CF and LDR were above the sensor detection limit of 5kg. The horses had significantly higher cortisol concentrations directly after LDR compared to LF. In addition, the horses showed more distinctive head movements, including head waving, during LDR. There were no significant treatment effects on HR and HRV. In conclusion, the results indicate that LDR may be more stressful to these horses during riding.
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Leliveld, L. M. C., Düpjan, S., Tuchscherer, A., & Puppe, B. (2016). Behavioural and physiological measures indicate subtle variations in the emotional valence of young pigs. Physiol. Behav., 157, 116–124.
Abstract: Abstract In the study of animal emotions, emotional valence has been found to be difficult to measure. Many studies of farm animals' emotions have therefore focussed on the identification of indicators of strong, mainly negative, emotions. However, subtle variations in emotional valence, such as those caused by rather moderate differences in husbandry conditions, may also affect animals' mood and welfare when such variations occur consistently. In this study, we investigated whether repeated moderate aversive or rewarding events could lead to measurable differences in emotional valence in young, weaned pigs. We conditioned 105 female pigs in a test arena to either a repeated startling procedure (sudden noises or appearances of objects) or a repeated rewarding procedure (applesauce, toy and straw) over 11 sessions. Control pigs were also regularly exposed to the same test arena but without conditioning. Before and after conditioning, we measured heart rate and its variability as well as the behavioural reactions of the subjects in the test arena, with a special focus on detailed acoustic analyses of their vocalisations. The behavioural and heart rate measures were analysed as changes compared to the baseline values before conditioning. A limited number of the putative indicators of emotional valence were affected by the conditioning. We found that the negatively conditioned pigs showed changes that were significantly different from those in control pigs, namely a decrease in locomotion and an increase in standing. The positively conditioned pigs, however, showed a stronger increase in heart rate and a smaller decrease in SDNN (a heart rate variability parameter indicating changes in autonomic regulation) compared to the controls. Compared to the negatively conditioned pigs, the positively conditioned pigs produced fewer vocalisations overall as well as fewer low-frequency grunts but more high-frequency grunts. The low-frequency grunts of the negatively conditioned pigs also showed lower frequency parameters (bandwidth, maximum frequency, 25% and 50% quartiles) compared to those of the positively conditioned pigs. In any of the statistically significant results, the conditioning accounted for 1.5–11.9% of variability in the outcome variable. Hence, we conclude that repeated moderate aversive and rewarding events have weak but measurable effects on some aspects of behaviour and physiology in young pigs, possibly indicating changes in emotional valence, which could ultimately affect their welfare. The combination of ethophysiological indicators, i.e., the concurrent examination of heart rate measures, behavioural responses and especially vocalisation patterns, as used in the current study, might be a useful way of examining subtle effects on emotional valence in further studies.
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Ramos, D., Reche-Junior, A., Fragoso, P. L., Palme, R., Yanasse, N. K., Gouvêa, V. R., et al. (2013). Are cats (Felis catus) from multi-cat households more stressed? Evidence from assessment of fecal glucocorticoid metabolite analysis. Physiol. Behav., 122, 72–75.
Abstract: Abstract Given the social and territorial features described in feral cats, it is commonly assumed that life in multi-cat households is stressful for domestic cats and suggested that cats kept as single pets are likely to have better welfare. On the other hand, it has been hypothesized that under high densities cats can organize themselves socially thus preventing stress when spatial dispersion is unavailable. This study was aimed at comparing the general arousal underpinning emotional distress in single housed cats and in cats from multi-cat households (2 and 3–4 cats) on the basis of fecal glucocorticoid metabolites (GCM) measured via enzyme immunoassay (EIA). GCM did not significantly vary as a function of living style (single, double or group-housing); highly stressed individuals were equally likely in the three groups. Young cats in multi-cat households had lower GCM, and overall cats that tolerate (as opposed to dislike) petting by the owners tended to have higher GCM levels. Other environmental aspects within cat houses (e.g. relationship with humans, resource availability) may play a more important role in day to day feline arousal levels than the number of cats per se.
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Wolter, R., Pantel, N., Stefanski, V., Möstl, E., & Krueger, K. (2014). The role of an alpha animal in changing environmental conditions. Physiol. Behav., 133, 236–243.
Abstract: Abstract The maintenance and development of conservation areas by grazing of large herbivores, such as Przewalski's horses, is common practice. Several nature conservation areas house male bachelor groups of this species. When males are needed for breeding they are removed from the groups, often without considering group compositions and individual social positions. However, alpha animals are needed for ensuring group stability and decision making in potentially dangerous situations in several species. To investigate the role of the alpha male in a bachelor group, we observed the behaviour of five Przewalski's horse males during the enlargement of their enclosure. We analyzed the group's social structure and movement orders, as well as the animals' connectedness, activity budgets, and whether they moved with preferred group members and how factors such as social rank influenced the horses' behaviour. We also investigated the excretion of glucocorticoid metabolites (GCM) via faeces of the horses while exploring a new area as a parameter of glucocorticoid production. Our results show that the alpha male is important for a bachelor group in changing environmental conditions. The alpha male had the highest level of connectedness within the group. When exploring the new environment, its position in the group changed from previously being the last to being the first. Furthermore the whole group behaviour changed when exploring the new area. The stallions showed reduced resting behavior, increased feeding and did not stay close to each other. We found that the excretion of glucocorticoid metabolites of most horses rose only marginally during the first days on the new area while only the alpha male showed a significant increased amount of glucocorticoid production during the first day of the enclosure enlargement.
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König von Borstel, U., Euent, S., Graf, P., König, S., & Gauly, M. (2011). Equine behaviour and heart rate in temperament tests with or without rider or handler. Physiol. Behav., 104(3), 454–463.
Abstract: The aim of the present study was to compare horses' heart rate (HR), heart rate variability (RMSSD, pNN50) and behaviour in the same temperament test when being ridden, led, and released free. Behavioural measurements included scores and linear measurements for reactivity (R), activity (A), time to calm down (T) and emotionality (E), recorded during the approach (1) and/or during confrontation with the stimulus (2). Sixty-five horses were each confronted 3 times (1 ridden, 1 led, 1 free running in balanced order) with 3 novel and/or sudden stimuli. Mixed model analysis indicated that leading resulted in the lowest (P < 0.05 throughout) reactions as measured by A1, A2, E1, E2, R2, and pNN50 while riding produced the strongest (A1, T2, HR, RMSSD, pNN50) or medium (E1, E2, R2) reactions. Free running resulted either in the strongest (A2, E1, E2, R2) or in the lowest (A1, T2, HR, RMSSD, pNN50) reactions. The repeatability across tests for HR (0.57), but not for RMSSD (0.23) or pNN50 (0.25) was higher than for any behavioural measurement: the latter ranged from values below 0.10 (A1, A2, T2) to values between 0.30 and 0.45 (E1, E2, R2). Overall, the results show that a rider or handler influences, but not completely masks, the horses' intrinsic behaviour in a temperament test, and this influence appeared to be stronger on behavioural variables and heart rate variability than on the horses' heart rates. Taking both practical considerations and repeatabilities into account, reactivity appears to be the most valuable parameter. Emotionality and heart rate can also yield valid results reflecting additional dimensions of temperament although their practical relevance may be less obvious. If a combination of observed variables is chosen with care, a valid assessment of a horse's temperament may be possible in all types of tests. However, in practice, tests that resemble the practical circumstances most closely, i.e. testing riding horses under a rider, should be chosen.
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Forkman, B., Boissy, A., Meunier-Salaün, M. - C., Canali, E., & Jones, R. B. (2007). A critical review of fear tests used on cattle, pigs, sheep, poultry and horses. Physiol. Behav., 92(3), 340–374.
Abstract: FORKMAN, B., A., BOISSY, M.-C., SALAUN, E., CANALI, AND R.B., JONES. A critical review of fear tests used on cattle, pigs, sheep, poultry and horses. PHYSIOL. BEHAV. 000-000, 2007. Fear is arguably the most commonly investigated emotion in domestic animals. In the current review we attempt to establish the level of repeatability and validity found for fear tests used on cattle, pigs, sheep and goats, poultry and horses. We focus the review on the three most common types of fear tests: the arena test (open field), the novel object test, and the restraint test. For some tests, e.g. tonic immobility in poultry, there is a good and broad literature on factors that affect the outcome of the test, the validity of the test and its age dependency. However, there are comparatively few of these well defined and validated tests and what is especially missing for most tests is information on the robustness, i.e., what aspects can be changed without affecting the validity of the tests. The relative absence of standardized tests hampers the development of applied ethology as a science.
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De Boyer Des Roches, A., Richard-Yris, M. - A., Henry, S., Ezzaouia, M., & Hausberger, M. (2008). Laterality and emotions: visual laterality in the domestic horse (Equus caballus) differs with objects' emotional value. Physiol. Behav., 94(3), 487–490.
Abstract: Lateralization of emotions has received great attention in the last decades, both in humans and animals, but little interest has been given to side bias in perceptual processing. Here, we investigated the influence of the emotional valence of stimuli on visual and olfactory explorations by horses, a large mammalian species with two large monocular visual fields and almost complete decussation of optic fibres. We confronted 38 Arab mares to three objects with either a positive, negative or neutral emotional valence (novel object). The results revealed a gradient of exploration of the 3 objects according to their emotional value and a clear asymmetry in visual exploration. When exploring the novel object, mares used preferentially their right eyes, while they showed a slight tendency to use their left eyes for the negative object. No asymmetry was evidenced for the object with the positive valence. A trend for an asymmetry in olfactory investigation was also observed. Our data confirm the role of the left hemisphere in assessing novelty in horses like in many vertebrate species and the possible role of the right hemisphere in processing negative emotional responses. Our findings also suggest the importance of both hemispheres in the processing positive emotions. This study is, to our knowledge, the first to demonstrate clearly that the emotional valence of a stimulus induces a specific visual lateralization pattern.
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